Vol. 11(9), pp. 127-142, October-December 2019 DOI: 10.5897/JENE2019.0799 Article Number: A3C45E862300 ISSN 2006-9847 Copyright © 2019 Author(s) retain the copyright of this article Journal of Ecology and The Natural Environment http://www.academicjournals.org/JENE

Full Length Research Paper

Forest land use and native trees diversity conservation in Togolese mega hotspot, Upper Guinean, West Africa

Kossi Adjossou1*, Fifonsi Ayélé Dangbo1, Tossimidé Houngbedji3, Komla Elikplim Abotsi1, Donko Koudzo Koda2, Atsu Kudzo Guelly2 and Kouami Kokou1

1Laboratoire de Recherche Forestière, Université de Lomé, Togo. 2Laboratoire de botanique et écologie végétale, Université de Lomé, Togo. 3Institut Togolais de Recherche Agronomique, Togo.

Received 14 September, 2019; Accepted 22 October, 2019

In tropics, species diversity in agricultural systems is often assessed without distinction between native and exotic species. However, the conservation value of an ecosystem depends on its richness in native species. This study was conducted to determine the conservation value of agricultural systems in Togo megahotspot, one of the species-rich sites in Upper Guinean. Specifically, the study compares fallow systems (FS), coffee systems (COFS), cocoa systems (COCS) to forest relics (FR) on the one hand, and on the other hand the agricultural systems (FS, COFS, COCS) between them base on natives tree species diversity and composition. Sites have been selected to represent different forest lands use types. Plots (n = 233) of different sizes (400, 500 and 625 m²) were used for data collection. In each plot, all living trees (DBH ≥ 10 cm) were counted. Rarefaction, generalized linear models (GLM) and ecological distance approach were used to standardized and compare the data. A total of 183 species were recorded, of which 42% were absent from the agricultural plots. Difference in diversity index were significant between the FR and agricultural systems (p<0.001), but not between agricultural systems (p=0.23). Guineo-Congolian climax and endemic species were seriously under threat. The study poses a real problem of regeneration dynamic of these species in human-dominated landscapes that requires further specific work.

Key words: Rainforests, agricultural practices, biodiversity conservation, Reducing Emissions from Deforestation and Forest Degradation (REDD+), Togo, Upper Guinean.

INTRODUCTION

Agroforestry is generally defined as association of trees especially traditional ones, play an important role as with crops and livestock. There are several agroforestry refuges of tropical biodiversity by providing habitat for practices; including shade trees with perennial crops many species of plants and animals in human-dominated such as coffee and cocoa. Several studies on tropical landscapes (Cicuzza et al., 2011; Cassano et al., 2012; ecosystems have shown that agroforestry systems, Maas et al., 2015; Gras et al., 2016). Thus, it is widely

*Corresponding author. Email: [email protected].

Author(s) agree that this article remain permanently open access under the terms of the Creative Commons Attribution License 4.0 International License 128 J. Ecol. Nat. Environ.

accepted that agroforestry is the only alternative to meet addition, the few studies carried out on the conservation the ecological and economic challenges posed by slash- value of agroforestry systems in Africa have taken into and-burn agriculture practices (Fischer and Vasseur, account exotic species, that is, species richness and not 2000). These ideas have contributed immensely to the forest species richness (Aerts et al., 2011). In Cameroon, growth of agroforestry projects across the tropics in the for example, cocoa systems are rich with 206 species of recent decades. Some of these studies have also trees while this richness included one third of exotic highlighted the importance of these systems for carbon species with densities which double those of native sequestration (Schroth and Harvey, 2007). Today, forest species (Sonwa et al., 2007). The disadvantage is that, policies of many counties aim at increasing forest areas exotic species can greatly influence diversity index whose through the promotion of agroforestry systems for carbon calculations are often based on abundance data, with as and biodiversity. The conservation of this latest consequence the overestimation of the conservation (biodiversity) is a critical component of the development value of these agroforestry systems. Otherwise, human and implementation of UN-led Reducing Emissions from land use causes major changes in species abundance Deforestation and Forest Degradation (REDD+) and composition, yet native and exotic species can (Bustamante et al., 2016). However, in recent decades, exhibit different responses to land use change. Native there have been major changes in agroforestry practices species populations generally decline in human‐impacted in the coffee-cocoa sector around the world because of habitats while exotic species often benefit (Jesse et al., the declining productivity due to tree age, soil poverty and 2018). Exotic species are often generalist species or increased disease. In West Africa, facing declining species that adapt easily to human disturbance. It is often productivity, traditional cocoa and coffee systems had those taxa that are of the highest priority for conservation, been the subject of several managements by producers such as regional forest endemics, that are the most to make them more productive. These included the extinction prone in modified tropical landscapes (Gardner rehabilitation and replanting of fields, the use of Upper et al., 2009). For these reasons, ecologists think that a Amazon hybrid cocoa without shade, intensification by better indicator of the conservation value of agroforest the planting of additional trees and fruit trees, and the systems would be the forest species richness. Thus, to conversion of secondary forest and fallow areas to estimate accurately the value of modified landscapes for agroforestry and monoculture plantations (Wessel and conserving regional forest biodiversity we need to know Quist-Wessel, 2015). These changes in agroforestry the proportion of species that inhabit human-modified management practices and their potential impact on systems that were also inhabitants of the original forest tropical biodiversity raise today the question of the landscape (Gardner et al., 2009). conservation value of agroforestry systems that requires The Upper Guinean forests are among the 25 most research, especially in Africa where the conservation important hotspots in the world (Poorter et al., 2004). value of agroforestry systems are poorly evaluated They are estimated to contain 2,800 forests plant species compared to Asia or Latine America (Fischer and among which 650 are endemics, and about 400 are Vasseur, 2000; Paruelo et al., 2001; Siebert, 2002; Rolim considered rare. Within Upper Guinean, three areas of and Chiarello, 2004; Ashley et al., 2006; Harvey et al., high diversity are distinguished and need protection and 2007; Schroth and Harvey, 2007b; Philpott et al., 2008; attention (Poorter et al., 2004). Togo megahotspot is one Chazdon et al., 2009; Gardner et al., 2009; Wanger et al., of the three areas of high diversity in Upper Guinean that 2009; Feeley and Silman, 2010; Hoehn et al., 2010; owes its immense biodiversity to its past climate but also Williams Guillén and Perfecto, 2010; Cicuzza et al., 2011; to the current ecological conditions of humidities and fogs Tscharntke et al., 2011; Cassano et al., 2012; Pelletier et which contributed to the installation of a great diversity of al., 2012; Dawson et al., 2013; Bustamante et al., 2014; forest communities (Akpagana, 1989). These forests of Maas et al., 2015; Sariyildiz et al., 2015; Sharma and the mountains of Togo have been assigned to agriculture, Vetaas, 2015; Valencia et al., 2015; Barsoum et al., particularly for shifting, coffee and cocoa cultivation, 2016; Gasperini et al., 2016; Gras et al., 2016; ; Farah et which has led to repeated clearings. The conservation of al., 2017; Almazán-Núñez et al., 2018; Benítez-Badillo et biodiversity in this area largely depends of a deeper al., 2018; Jesse et al., 2018; Rodrãguez-Echeverry et al., comprehension of the impact of forest land-use types on 2018; Santos-Heredia et al., 2018). Few studies in Africa native species diversity. However, this information is now compare agriculture systems to natural forests and absent today on this area. Togo Mountains vegetation each other to assess their conservation value based on was the subject of several works (Aké Assi, 1971; Brunel, forest species richness (Eilu et al., 2003; Augusseau et 1977, 1978; Akpagana, 1989, 1992; Guelly, 1994; al., 2006; Bobo et al., 2006; Norris et al., 2010; Pinard et Adjossou, 2004, 2009; Kokou et al., 2008; Adjossou and al., 2013). In Africa, several studies have been conducted Kokou, 2009;). Recent works have been focused on on agroforestry systems for coffee and cocoa. However, agroforestry systems in the area (Adden et al., 2016, these studies have rather addressed issues related to the 2018; Koda et al., 2016; Adden et al.). Nevertheless, decline in productivity and the socio-economic aspects of there is an important lack of information on conservation these systems (Wessel and Quist-Wessel, 2015). In value of forests land-use types of this area. The present Adjossou et al. 129

study is significant in providing clear answers to introduced into the forest zone. Until 1989, most of Togo's questions regarding the biodiversity value of different rainforests were converted into high-yielding coffee and cocoa forest land-use types in order to help policymakers and fields through massive fertilizer use. Conversion was possible thanks to the introduction and use of chainsaws. Since 1990, managers to conserve and manage the native trees producers could no longer buy fertilizer because of falling purchase species in most species-rich site of Togo. Togo has prices following the fall in world prices causing a drastic drop in signed the World Bank forest carbon partnership. This productivity. In response to the declining productivity of traditional study will guide the policies in the strategic choices and coffee-cocoa systems in the 1990s, farmers have chosen the implementations of REDD + projects in order to conserve management method based on intensification by the planting of additional trees and fruit trees. Currently, the use of Upper Amazon and manage the native tree species in Togolese hybrid cocoa without shade is emerging in the area. The meagahotspot. The aim of this study was to evaluate the conversions of secondary forest and fallow areas to agroforestry conservation value of agricultural systems in the Togo and monoculture plantations are also underway in Togo's forest Mgahotspot. Specifically, the study compares fallow zone. What remains of these forests today is very fragmented and systems (FS), coffee systems (COFS), cocoa systems primarily localized in the zones difficult to reach, and along the (COCS) to forest relics (FR) on the one hand, and on the rivers. Because of a high demographic growth rate in Togo (3.3% annually), the forest remnants are also affected, contrary to those other hand the agricultural systems (FS, COFS, COCS) along the rivers which are still relatively saved by the local between them base on natives tree species diversity and populations, partly for traditional reasons (Adjossou, 2009). composition.

Sampling methods

METHODS The forest land-use types were identified and inventoried through reconnaissance survey in the field, involving local populations but Study area also on the basis of available historical vegetation maps which distinguishes forest areas from savannas. This approach was used Forest growth conditions because of the unavailability of accurate geographic information on each forest land- use types in the study area. These The studied area (6°15‟ N - 8°20‟ N; 0°30‟ E -1° E) covers 6,441 reconnaissance survey and data collections were carried out Km² in the southern part of the Atakora mountains, south-west of between 2007 and 2012. Sites were selected to represent study Togo, on the border between Togo and Ghana called Togo land-use types. In each site, sampling points were selected Mountains or Togo highlands or Ecological Zone IV (Figure 1). It is systematically to represent the physiognomy of different land-use. adjacent to Dahomey Gap, an extension of the woodland savannas Plots (n=223) of different sizes (400, 500 and 625 m²) of the Sahel to the Gulf of Guinea, which separates the Upper corresponding to an area of 16 ha were used to study the native Guinea forests from the rest of the African rainforests (Poorter et plant communities under different forest land use: FS (n= 41; 3.35 al., 2004). The subhumid mountainous area of Togo owes its ha), COFS (n=43; 3.12 ha), COCS (n=72; 5.13 ha), FR (n=67, 4.41 immense biodiversity not only to its past climate but also to the ha). Plots 0.04 ha in area (20 m × 20 m) were used to study the current ecological conditions of humidities and fogs which native plant communities under forests relic land-use type. But, if contributed to the installation of a great diversity of forest the habitat of a plant community was long and narrow such as for communities. The areas are mainly covered with deciduous forest ones located near a stream, 10 m × 50 m plots were used (Kokou, (Akpagana, 1989) intersected with Guinean savannas (Guelly, 1998; Natta, 2003). Under coffee and cocoa systems, plots 0.0625 1994). Six types of upland semi-deciduous forest were ha in area (25 m × 25 m) were used. Under FS, plots 0.04 ha in distinguished: (1) Forest with Sterculiaceae and Sapotaceae; (2) area (20 m x 20 m) and 0.0625 ha (25 m × 25 m) were used. In Forest with dominant Celtis mildbraedii; (3) Forest with dominant these different plots, all living natives‟ trees with a DBH ≥10 cm Terminalia superba; (4) Forest with dominant Ricinodendron (diameter at 1.3 m height) were counted. heudelotii; (5) Forest with Meliaceae and Moraceae; (6) Forests with Parinari excelsa (Akpagana, 1989). Adjossou et al (submitted) found that the forest vegetation of the site is rather a mixture of Data analysis Guineo-Congolese forest types. The climate in this area is of Guinea Mountain type (Papadakis, 1966) characterized by a long Species richness and diversity rainy season (8-10 months). The mean annual temperatures range from 21 to 25°C and the total annual rainfall ranges from 1250 to To compare species richness and diversity between land-use types, 1900 mm.. Landforms are diverse and complex. The main geologic species accumulation curves, gamma, alpha, beta, and Shannon component is of the late Precambrian (Hall and Swaine, 1976). The diversity were used. According to Gotelli and Colwell (2001), if the main edaphic component consists of schist. sampling methods are not identical, different kinds of species may be over or under-represented in different samples. To solve the problem of inequality samples size in this study (400, 500 and 625 History of forest land-use m ²), different data standardization techniques were used. Firstly, richness of different plot sizes were standardized to the same 400 Three main forestry uses can be distinguished in the mountains of m² plot following Vandermeer et al. (2000). Secondly, samples- Togo, which are shifting, coffee and cocoa cultivation. Coffee and based species accumulation curves were used to standardize and cocoa were introduced into the humid forest zone of Togo around compare total species richness between the four forest land-use 1945. The varieties introduced at the time were Coffea canefera types based on a sampling area of 400 m². This rarefaction method (GNAWLUI) Coffea robusta (ROBOSCA) for coffee and Theobroma is especially useful when comparing species richness between sp (TETEKOSSI) for cocoa. These traditional varieties were grown subsets of different sizes (Kindt and Coe, 2005) because it allows in rainforests understory and did not pose a real threat to plant comparing the same number of plots for each subset. Species biodiversity. From 1975, sun-loving hybrids (Agric varieties) were accumulation curves of each forest land use were calculated from 130 J. Ecol. Nat. Environ.

Figure 1. Map showing the location of the study sites in Togo megahotspot (Ecological zone IV). Adjossou et al. 131

Table 1. A comparison of total observe species number, number of unique species, number of climax species, mean species richness, mean Shannon diversity index; mean abundance of native trees species (DBH> 10 cm) on different forest land used types in Togo megahopspot mixed evergreen/semidecidous forest zone

Parameter FR COCS COFS FS Total area sampled (ha) 4.41 5.13 3.12 3.35 Gamma diversity (Area sampled) 165 72 67 65 Number of unique species (Area sampled) 77 8 4 3 Alpha diversity per plot (400m²) 13.76 (SD=5.66) 4.15 (SD=3.11) 4.72 (SD=2.75) 4.88 (SD=4.43) Beta diversity per plot (400m²) 10.99 16.33 13.19 12.32 Mean Shannon diversity index per plot (400 m²) 2.54 (SD=0.42) 1.15 (SD=0.79) 1.40 (SD=0.59) 1.28 (SD=0.82) Mean abundance per plot (400m²) 10.07 (SD=5.52) 2.25 (SD=1.64) 2.55 (SD=1.30) 2.38 (SD=2.32)

FR: Forest Relics; COCS: Coco System; COFS: Coffee System; FS: Follow Systems.

the cumulative number of species based on the number of species following Misra (1968). All the data supporting this paper are present in the plot sample using bootstrap‟s randomization (200 available at times) re-sampling techniques. https://data.mendeley.com/datasets/kymphvj2cj/draft?a=52c6ccfd- Gamma, alpha and beta diversity were used as the basic fdb2-4c2e-8a05-daaf46d2f10d diversity index (Whittaker, 1972). Gamma diversity is the landscape-level diversity estimated as the total number of species or total richness across plots. Alpha diversity is calculated as the RESULTS average species richness per plot. Beta diversity is a measure of heterogeneity in the data or habitats diversity (McCune et al., Species richness and diversity 2002), defined as: βw = (Sc/S)-1, where βw is the beta diversity, Sc is the number of species in the whole data set and S is the average A total of 183 native tree species (Appendix S1) species richness in the sample units. The one is subtracted to make zero beta diversity correspond to zero variation in species comprising 2485 stems were recorded. They were abundance. If βw = 0, then all sample units have all of the species. divided into 165 (89% of the total species recorded) The larger the value of βw, the more heterogeneous the data set is. species and 1759 (61% of the total stems recorded) As a rule of thumb in that context, values of βw < 1 are rather low stems, 72 (39%) and 420 (15%), 67 (36%) and 330 and βw > 5 can be considered high. The maximum value of βw is (11%) and 65 (35%) and 366 (12%) on FR, COCS, Sc-1. The maximum value is obtained when no species are shared among sample units. Shannon–Wiener index (H) was computed COFS and the FS, respectively (Table 1). It is worth following Magurran (1988): noting that the largest area was sampled in the Cocoa System (COCS). However, total species richness in this H = − Σ (pi) Log2 (pi) system, similar to that of Coffee System (COFS) and Fallow system (FS), remains the lowest compared to Where pi=Ni/N is the proportional abundance of species i and Log2 forest relics. is the base of the logarithm. Comparison of the accumulation curves between the Generalized linear models (GLM) was used with binomial (link= four forest land use types, based on the same number of logit) variance functions to standardize and compare alpha and plots (40 plots of 400 m² for each type of land use), Shannon index between forests land use. This technique offer the best approach because it does not require that sample sizes to be showed that FR was the richest in observed species (143 equal, as required to compare the total number of species (Kindt species), followed by COFS and FS (63 species each) and Coe, 2005). Analysis of variance is carried out to examine and COCS (57) (Figure 2). Thus, the species richness of whether the data are consistent with a simpler („null‟) model in agricultural systems accounted only for 40 to 44% of which there are no differences. forest relics. Alpha and Shannon diversity were three times higher in the RF than in the three agricultural forest land use types (Table 1). However, the beta diversity Floristic similarity and species abundance results showed a high value in the COCS (16.33; SD=3.10), followed by COFS (13.19, SD=2.74), FS To assess differences in species composition among forest land use types, similarity method based on abundance of species was (12.32, SD=4), and FR (10.99; SD=5.66). Difference in performed using Bray-Curtis ecological distance. Bray-Curtis alpha (LR Chisq = 135.38; P<0.001) and Shannon distance was chosen because it is the most widely used abundance diversity (LR Chisq = 142.61; P<0.001) were significant based measure, due to its strong relationship to ecological distance between the FR and the three agricultural systems. But, under varying conditions (Bray and Curtis, 1957). To diminish the there is no significant difference in alpha (LR Chisq = influence of the dominant species, logarithmic transformation technique has been applied to the species matrix. To assess haw 1.41 P=0.23) and Shannon diversity (LR Chisq = 2.5 species abundance varied among forest land use types species P=0.11) between the three agricultural systems indicating abundance as the number of individuals per ha was calculated that the studied agroforestry systems for coffee and 132 J. Ecol. Nat. Environ.

Figure 2. Samples-based species accumulation curves of the four forest land use types in the Togo Megahotspot. FR: Forest Relics; COCS: Coco System; COFS: Coffee System; FS: Follow Systems. All curves are based on a sampling area of 400 m².

cocoa system have the same conservation value as africana (21.35), Pycnanthus angolensis (13.60), fallows systems. Sterculia tragacantha, Tabernaemontana pachysiphon, Pterocarpus mildbraedii, Cola gigantea, Albizia adianthifolia, and Antiaris toxicaria subsp. Africana. In the Floristic similarity and species abundance COCS, Milicia excelsa (9.55 ind/ha) was the abundant distribution species with the highest density, followed by Terminalia superba (5.06), Funtumia africana (4.28), Ficus mucuso, The overall Bray-Curtis distance values between forest Sterculia tragacantha, Ficus sur, Monodora myristica, lands use types ranged from 0.39 to 0.60. Highest Bray- Khaya grandifoliola etc. In the COFS, Xylopia aethiopica Curtis distance (D=0.6) was observed between FR and (12.17 ind/ha) was the abundant species, followed by agricultural systems and less Bray-Curtis distance (D = Milicia excelsa (9. 29), Albizia adianthifolia (8.65), Albizia 0.39) was observed between the three agricultural zygia (7.05), Khaya grandifoliola, Terminalia superba, systems (Table 2). FR shared 32 (17.3% of total species) Anthocleista djalonensis, Morinda lucida, Monodora species with the three agricultural systems, whereas 77 myristica, Aubrevillea kerstingii etc. (Appendix S1) (42% of total species) species occurred on FR site only. The majority of species (78%) have low abundance Only 8 (4%), 4 (2%) and 3 (2%) species were unique to (density <5 ind/ha) and could be considered rare. Of COCS, COFS and FS site, respectively. these, 57% were endemic to Guneo-Congolese Abundance distribution patterns showed that species rainforests (Afzelia bella, Cordia platythyrsa, Drypetes individual abundance varied between forest lands uses aylmeri, Entandrophragma cylindricum, Erythrina vogelii, types. The most abundant species by number of stems Gymnostemon zaizou, Homalium letestui, Parkia bicolor, recorded in the FR was Pseudospondias microcarpa (30 Pterygota bequaertii etc.), 21% of the transition species ind /ha); followed by Celtis mildbraedii (22.22), Funtumia between Guneo-Congolese rainforests and dry Adjossou et al. 133

Table 2. Distance matrix between forest land use types in Togo megahopspot mixed evergreen/semidecidous forest zone using the Bray- Curtis , Kulczynski and Gower distances

Distance FR-COCS FR-COFS FR-FS COCS-COFS COCS-FS COFS-FS Bray- Curtis distance 0.58 0.58 0.56 0.39 0.44 0.40 Kulczynski distance 0.56 0.56 0.53 0.39 0.44 0.40 Gower distance 0.76 0.75 0.72 0.30 0.35 0.31

FR: Forest Relics; COCS: Coco System; COFS: Coffee System; FS: Follow Systems.

vegetation (Afzelia Africana, Anogeissus leiocarpus, in this results (SD) (Table 1). However, this heterogeneity Cassia sieberiana, Eriocoelum kerstingii, Malacantha was more pronounced in cocoa (beta = 16) and coffee alnifolia, Manilkara multinervis etc.) (Appendix S1), (13) systems. This could be related to the different management schemes. Very few studies have compared the native species DISCUSSION richness and composition between agricultural systems in tropics. In general, both plant and animal diversity within Results showed a high species richness and diversity in cocoa agroforests is greater than those of other the FR compared to the three agricultural systems agricultural land uses, but lower than in the original forest (COCS, COFS and FS). Very few studies in the tropics habitats (Harvey et al., 2007). It has been shown that have reported the impacts of cocoa and coffee fallows are less diversified than cocoa systems (Trimble agroforests practices on the richness and diversity of and Van Aarde, 2014). However, based on the native native tree species compared to those that included species, this study has shown that agricultural systems exotic species in the calculation of diversity index, or (COCS, COFS and FS), in Togolese megahotspot, have focused on the wildlife richness of these systems (Harvey similar species diversity and composition, hence similar and González Villalobos, 2007). Rolim and Chiarello conservation values. This could be justified by changes in (2004) have been reported that the cocoa agroforestry the management practices of coffee-cocoa agroforestry (cabruca forests) systems in southeastern Brazil were systems which are causing their change close to FS. less diverse and less dense than secondary or primary These changes affect native species identity and forests of the region. In Densu Basin in Ghana, Attua abundance distribution across the study area and the (2003) (cited by Norris et al., 2010) showed that the plant types of forest land use. At the scale of the study area, community of existing pockets of forest are more species very few species were abundant (22%) (Table 4), while diverse than the vegetation associated with the cocoa or the majority could be considered rare (78%). In addition, food crop farms. 42% of species recorded were absent from agricultural Findings showed that native‟s tree species richness systems; some species have low densities on agricultural was reduced by 70, 66 and 65% and Shannon index by systems compared to their density in FR. Moreover, it 55, 45 and 50% in COCS, COFS and FS, respectively was noted that: compared to FR. The reduction of regional species richness by coffee systems in this study (66%) can be 1. Species abundance varied according to the types of considered high compared to that observed in similar practices. For example Milicia excelsa and Terminalia systems in Ethiopia (25%) (Aerts et al., 2011). This superba were abundant in the COCS and COFS, while difference could be due to the fact that the two studies Albizia adianthifolia and Albizia zygia in the COFS and did not use the same methods. In this study, trees of FS. Ficus mucuso and Funtumia africana were abundant DBH ≥10 cm were considered, exotic species were not in the COCS, Xylopia aethiopica in the COFS and taken into account while Aerts et al. (2011) considered Macaranga barteri in the FS (Table 4); species ≥ 2 m tall and exotic species. This observations, 2. The most abundant species at the scale of the study with regard to Shannon diversity Index, are in agreement area included species with high economic values with those of Rodriguez-Echeverry et al. (2018) who have belonging to the categories of timber species (Khaya shown that the land-use change decreased Shannon grandifoliola, Milicia excelsa, Terminalia superba etc.), diversity index in the Chilean temperate forests medicinal species (Alstonia boonei, Anthocleista (Rodrãguez-Echeverry et al., 2018). djalonensis and Spathodea campanulata) and spice The results showed a high beta diversity index in the species (Monodora myristica and Xylopia aethiopica) and four forest land use types (> 5) showing a great multipurpose species (fertilization, soil restoration, heterogeneity in the data which could be explained by the shading, firewood, fruits, tools, etc.) and pioneer species variability of species abundance within and between like: Albizia adianthifolia, Albizia adianthifolia, Funtumia forest land use types, justifying the differences observed Africana, Ficus exasperata, Ficus mucuso, Ficus sur, 134 J. Ecol. Nat. Environ.

Table 3. Pioneer and farmers' preferred dominated tree species in Togo megahopspot mixed evergreen/semidecidous forest zone. The numbers represent the density per ha.

Species Families FR COCS COFS FS Total Milicia excelsa (Welw.) C.C. Berg. Moraceae 6.12 9.55 9.29 2.38 27.35 Funtumia africana (Benth.) Stapf Apocynaceae 21.31 4.28 0.96 0.29 26.86 Albizia adianthifolia (Schum.) W. F. Wright Fabaceae 7.48 1.16 8.65 6.86 24.17 Albizia zygia (DC.) J.F. Macbr. Fabaceae 2.04 1.75 7.05 11.64 22.48 Pycnanthus angolensis (Welw.) Warb. Myristicaceae 13.6 1.55 2.24 4.47 21.88 Sterculia tragacantha Lindl. Sterculiaceae 12.01 3.89 1.6 3.88 21.39 Khaya grandifoliola DC. Meliaceae 7.02 2.72 4.8 3.28 17.84 Ficus sur Forssk Moraceae 4.98 3.89 1.6 6.86 17.35 Morinda lucida Benth. Rubiaceae 4.08 2.72 4.16 6.26 17.24 Macaranga barteri Müll. Arg. Euphorbiaceae 5.44 0.19 1.6 8.95 16.19 Terminalia superba Engl. & Diels Combretaceae 4.76 5.06 4.48 0.29 14.61 Cola gigantea var. glabrescens Brenan & Keay Sterculiaceae 8.39 1.55 0.96 3.28 14.19 Xylopia aethiopica (Dural) A. Rich. Annonaceae 0.22 0.77 12.17 0.29 13.48 Monodora myristica (Gaertn.) Dunal Annonaceae 5.66 3.31 3.84 0.29 13.12 Ficus exasperata Vahl. Moraceae 4.08 1.36 1.6 5.37 12.42 Ficus mucuso Welw. ex Ficalho Moraceae 5.21 4.09 0.64 1.79 11.74 Antiaris africana Engl. Moraceae 7.48 0.97 0.96 2.08 11.5 Aubrevillea kerstingii (Harms) Fabaceae 6.57 0.19 3.2 0.59 10.57 Anthocleista djalonensis A. Chev. Loganiaceae 2.49 1.75 4.48 1.79 10.52 Alstonia boonei De Wild. Apocynaceae 4.53 1.55 0.32 3.88 10.29 Ceiba pentandra (Linn.) Gaerth. Bombacaceae 4.98 1.94 0.32 2.38 9.64 Canarium schweinfurthii Engel. Burseraceae 6.12 0.19 0.32 1.49 8.13 Tetrorchidium didymostemon (Baill.) Pax & Hoffm. Euphorbiaceae 3.85 0.58 0.32 1.49 6.25 Holarrhena floribunda (G. DON.) Dur. & Schinz. Apocynaceae 2.26 1.36 1.6 0.59 5.83 Spathodea campanulata P. Beauv. Bignoniaceae 0.9 1.36 1.28 1.79 5.34 Harungana madagascariensis Lam. ex Poir. Guttiferae 0.9 0.38 2.24 1.49 5.03 Vitex doniana Sweet Verbenaceae 1.36 1.16 1.28 1.19 5 Ricinodendron heudelotii (Baill.) Pierre ex Pax Euphorbiaceae 1.36 1.94 0.32 0.29 3.92 Margaritaria discoidea (Baiil) Webster Phyllanthaceae 0.22 0.38 0.64 2.38 3.64

FR: Forest Relics; COCS: Coco System; COFS: Coffee System; FS: Follow Systems.

Harungana madagascariensis, Holarrhena floribunda, forest products. Immediate useful species were those Macaranga barteri, Margaritaria discoidea, Morinda plants that the farmers use for food, medicine, firewood, lucida, Tetrorchidium didymostemon etc.; among them, construction, or raw materials (Diemont et al., 2011). The there are several pioneer species (Table 3). results are also in line with those obtained by Adden et al. These distribution patterns were a reflection of the (2018) who studied the preference of trees by farmers in selection of trees by farmers. These results are the coffee and cocoa agroforestry systems in Togo. Their consistent with those of other authors. Indeed, works study showed that Togolese farmers prefer in their based on interviewed and tree inventories in coffee farms plantation species like Albizia spp., Khaya grandifoliola, and forest sites in La Sepultura Biosphere Reserve in Milicia excels and Terminalia superba. Chiapas, Mexico have showed that farmers are modifying These patterns suggested also that agricultural agroforests according to their knowledge and tree systems block the natural succession of certain species preferences, and that the resulting agroforest is lower in especially climax and endemic species of Guineo- tree diversity and dominated by pioneer and farmers' congolian. These results are in agreement with those of preferred tree species as compared to forests (Valencia other authors. In semi-forest coffee agroforestry system et al., 2015). In Agroforestry system in southern Mexico in Ethiopian Afromontane rainforest fragments, climax and Central America, more than 30 native tree species species of the rainforest were underrepresented (Aerts et are recognized and managed as potential facilitators of al., 2011). Waltert et al. (2011) assessed conservation forest regeneration and direct human consumption of values in tropical land use systems and found that Adjossou et al. 135

Table 4. Some Guineo-Congolian endemic and climax endangered species in Togo megahopspot mixed evergreen/semidecidous forest zone.

Species Family FR COCS COFS FS Total Parinari glabra Oliv. Chrysobalanaceae 3.4 - - - 3.4 Aningeria altissima (A. Chev.) Aubrév. et Pellegr. Sapotaceae 3.17 - - - 3.17 Piptadeniastrum africanum (Hook. f.) Brenan Fabaceae 2.26 0.58 0.32 - 3.17 Polyscias fulva (Hiern) Harms Araliaceae 3.17 - - - 3.17 Hannoa klaineana Pierre et Engl. Simaroubaceae 0.68 0.19 1.6 0.59 3.07 Nesogordonia papaverifolia (A.Chev.) R. Capuron Sterculiaceae 2.72 - 0.32 - 3.04 Sterculia oblonga Mast. Sterculiaceae 2.72 - - 0.29 3.01 Bombax buonopozense P. Beauv. Bombacaceae 0.22 0.97 0.32 1.19 2.71 Zanthoxylum macrophylla Engl. Rutaceae 0.68 - 1.92 - 2.6 Holoptelea grandis (Hutch.) Mildbr. Ulmaceae 2.26 0.19 - - 2.46 Turraeanthus africanus Pellegr. Meliaceae 2.26 - - - 2.26 Pterygota macrocarpa K. Schum. Sterculiaceae 1.81 - 0.32 - 2.13 Distemonanthus benthamianus Baill. Fabaceae 0.9 0.19 0.96 - 2.06 Maesopsis eminii Engl. Rhamnaceae 2.04 - - - 2.04 Lovoa trichilioides Harms Meliaceae 1.58 - - - 1.58 Amphimas pterocarpoïdes Harms Fabaceae 0.22 0.38 0.64 0.29 1.55 Trichilia megalantha Harms Meliaceae 1.13 0.38 _ _ 1.52 Tetrapleura tetraptera (Schum.& Thonn.) Taub. Fabaceae 1.13 - 0.32 _ 1.45 Khaya anthotheca (Welw.) C.DC. Meliaceae 0.45 - 0.32 0.59 1.37 Afzelia bella Harms var.gracilior Keay Fabaceae 1.36 - - - 1.36 Morus mesozygia Stapf Moraceae 0.68 - 0.32 0.29 1.29 Klainedoxa gabonensis Pierre ex Engl. Irvingiaceae 0.68 0.38 - - 1.07 Sterculia rhinopetala K. Schum. Sterculiaceae 0.68 0.38 - - 1.07 Treculia africana Decne Moraceae 0.45 - - 0.59 1.05 Parinari excelsa Sabine Chrysobalanaceae 0.68 - 0.32 - 1 Gymnostemon zaizou Aubrév. & Pellegr. Simaroubaceae 0.9 - - - 0.9 Homalium letestui Pellegr Flacourtiaceae 0.9 - - - 0.9 Afrosersalisia afzelii (Engel.) A. Chev. Sapotaceae 0.45 - 0.32 - 0.77 Blighia welwitschii (Hiern) Radlk Sapindaceae 0.22 0.38 - - 0.61 Nauclea diderrichii (de Wild. & Th. Dur. Merril.) Rubiaceae _ 0.58 - - 0.58 Drypetes gilgiana(Pax)(Pax)&K.Hoffm. Euphorbiaceae 0.45 _ - - 0.45 Entandrophragma cylindricum (Welw.) C.DC. Meliaceae 0.45 _ - - 0.45 Stereospermum acuminatissimum K. Schum. Bignoniaceae 0.45 _ - - 0.45 Tarenna pavettoides (Harv.) Sim Rubiaceae 0.45 _ - - 0.45 Mansonia altissima A. Chevalier Sterculiaceae 0.22 0.19 - - 0.42 Cordia platythyrsa Baker Boraginaceae 0.22 - - - 0.22 Drypetes aframensis Hutch. Euphorbiaceae 0.22 - - - 0.22 Drypetes aylmeri Hutct .& Dalz. Euphorbiaceae 0.22 - - - 0.22 Drypetes leonensis Pax Euphorbiaceae 0.22 - - - 0.22 Erythrina vogelii Hook. f. Fabaceae 0.22 - - - 0.22 Irvingia robur Mildbr. Irvingiaceae 0.22 - - - 0.22 Parkia bicolor A.Chev. Fabaceae 0.22 - - - 0.22 Parkia filicoidea Welw. ex Oliv. Fabaceae 0.22 - - - 0.22 Pterygota bequaertii De Wild. Sterculiaceae 0.22 - - - 0.22 Rhodognaphalon brevicuspe (Sprague) Roberty Sterculiaceae 0.22 - - - 0.22 Symphonia globulifera Linn. f. Guttiferae 0.22 - - - 0.22 Erythrina mildbraedii Harms Fabaceae _ 0.19 - - 0.19

FR: Forest Relics; COCS: Coco System; COFS: Coffee System; FS: Follow Systems. The numbers represent the density per hectare. 136 J. Ecol. Nat. Environ.

agroforestry systems, even traditional, reduced by 91% availability. These measures are good except that endemic plant species and concluded that even maintaining or restoring floristically diverse and ecologically friendly agricultural matrices may be of much structurally complex shade canopies within cocoa lower value for tropical conservation than indicated by agroforests are at the origin of the problems that the mere biodiversity value. The results of this study confirm cocoa sector is facing today (disease, rot, yield those of Barlow et al. (2007) who quantified the reduction). This study showed that 78% of the native biodiversity value of tropical primary, secondary, and species of the Megahotspot of Togo are threatened. Of planting and found that almost 60% of plant species were these, 57% of climax and endemic species of the Guineo- only ever recorded in primary forest. Congolian region, 21% of the transition species between Finally, the study showed that some species have low Guneo-Congolese rainforests and dry vegetation. In density in both FR and agricultural systems. These terms of conservation, transitional species are not results indicated that agroforestry is not the only cause of endangered because of their wide distribution. They can species rarity in the area. The exploitation of the timber be found in other types of vegetation. But the climax and could explain the rarity of species. Pseudospondias endemic species of the Guineo-Congolian region deserve microcarpa was the most abundant species in FR special attention (Table 4). Measures will be taken to because it is not a timber species. ensure the renewal of these species in both FR and agricultural systems. This study poses a real problem of regeneration and succession of tropical rainforest species Implication for management and biodiversity in human-dominated landscapes. Of the characteristic conservation species, it is the anthropophilic and heliophilous species that colonize our agricultural and forest landscapes. Agroforestry systems are often presented as a refuge for Important ecological issues are required for the tropical biodiversity, only alternative to meet the conservation and sustainable management of biodiversity ecological and economic challenges posed by slash-and- in this area. This requires scientific and indigenous burn agriculture practices (Fischer and Vasseur, 2000). knowledge. Bhagwat et al. (2008) reviewed evidence from studies across the tropics where species richness and composition of agroforestry systems are compared with CONFLICTS OF INTEREST that of neighbouring forest reserves and concluded that agroforestry systems are high in species richness and The authors declare no conflict of interest. more similar to neighbouring forest reserves in species composition if (i) the forest land was fairly recently converted to agroforestry plantation; (ii) the management ACKNOWLEDGEMENTS was less intensive; and (iii) the canopy cover of native trees was high. In traditional systems where these The authors would like to express their appreciation to conditions are met, significant reductions in native the local people of rainforest zone of Togo for their species could have been observed. Waltert et al. (2011) reception and field assistance. This research was have shown that traditional agroforestry systems could sponsored by ITTO Fellowship (Ref. 071/03A), BES reduce more than 90% of native plant species. This Overseas Research Grant (Project RE. 449 / 491), IFS- reduction is not only related to the management practices Comintech Research Grant (Project D/4037-1; Project of these systems but also their effect on microhabitats. D/4037-2; ProjectD/4767 ) and AUF Fellowship. Tropical forest ecosystems consist of a large number of microhabitats, each of which supports a high diversity of specialized organisms. Many organisms depend upon REFERENCES these microhabitats for a critical stage in their life cycle. Management that removes or modifies any of these Adden AK, Adjonou K, Kokou K, Agbemanyale K (2016). Cocoa trees microhabitats can potentially cause critical changes in (Theobroma cacao linn.) agroforests replanting in Togo: What appropriate agroecosystems? 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Appendix S1

Appendix S1. Species abundance in Togo megahopspot mixed evergreen/semidecidous forest zone. The numbers represent the density per ha

Species Fam Succession Chie FR COCS COFS FS Total Pseudospondias microcarpa ( A. Rich.) Engl. Anacardiaceae Pioneer GC-SZ 29,93 _ _ 1,79 31,72 Milicia excelsa (Welw.) C.C. Berg. Moraceae Climax GC 6,12 9,55 9,29 2,38 27,35 Funtumia africana (Benth.) Stapf Apocynaceae Pioneer GC 21,31 4,28 0,96 0,29 26,86 Albizia adianthifolia (Schum.) W. F. Wright Fabaceae Pioneer GC 7,48 1,16 8,65 6,86 24,17 Albizia zygia (DC.) J.F. Macbr. Fabaceae Pioneer GC 2,04 1,75 7,05 11,64 22,48 Celtis mildbraedii Engl. Ulmaceae Climax GC 22,22 _ _ _ 22,22 Pycnanthus angolensis (Welw.) Warb. Myristicaceae Climax GC 13,6 1,55 2,24 4,47 21,88 Sterculia tragacantha Lindl. Sterculiaceae Pioneer GC-SZ 12,01 3,89 1,6 3,88 21,39 Khaya grandifoliola DC. Meliaceae Climax GC 7,02 2,72 4,8 3,28 17,84 Ficus sur Forssk Moraceae Pioneer GC-SZ 4,98 3,89 1,6 6,86 17,35 Morinda lucida Benth. Rubiaceae Pioneer GC-SZ 4,08 2,72 4,16 6,26 17,24 Macaranga barteri Müll. Arg. Euphorbiaceae Pioneer GC 5,44 0,19 1,6 8,95 16,19 Terminalia superba Engl. & Diels Combretaceae Climax GC 4,76 5,06 4,48 0,29 14,61 Cola gigantea var. glabrescens Brenan & Keay Sterculiaceae Climax GC 8,39 1,55 0,96 3,28 14,19 Xylopia aethiopica (Dural) A. Rich. Annonaceae Pioneer GC-SZ 0,22 0,77 12,17 0,29 13,48 Monodora myristica (Gaertn.) Dunal Annonaceae Climax GC 5,66 3,31 3,84 0,29 13,12 Ficus exasperata Vahl. Moraceae Pioneer GC-SZ 4,08 1,36 1,6 5,37 12,42 Ficus mucuso Welw. ex Ficalho Moraceae Pioneer GC 5,21 4,09 0,64 1,79 11,74 Antiaris africana Engl. Moraceae Pioneer GC-SZ 7,48 0,97 0,96 2,08 11,5 Aubrevillea kerstingii (Harms) Fabaceae Climax GC 6,57 0,19 3,2 0,59 10,57 Anthocleista djalonensis A. Chev. Loganiaceae Pioneer GC-SZ 2,49 1,75 4,48 1,79 10,52 Alstonia boonei De Wild. Apocynaceae Climax GC 4,53 1,55 0,32 3,88 10,29 Ceiba pentandra (Linn.) Gaerth. Bombacaceae Pioneer GC-SZ 4,98 1,94 0,32 2,38 9,64 Cleistopholis patens (Benth.) Engl. & Diels Annonaceae Pioneer GC 6,57 2,14 _ 0,59 9,31 Tabernaemontana pachysiphon Stapf Apocynaceae Climax GC 9,07 _ _ _ 9,07 Pterocarpus mildbraedei Harms Fabaceae Climax GC 8,61 _ _ 0,29 8,91 Cola millenii K. Schum. Sterculiaceae Climax GC 7,48 _ 0,32 0,59 8,4 Canarium schweinfurthii Engel. Burseraceae Climax GC 6,12 0,19 0,32 1,49 8,13 Trilepisium madagascariense DC. Moraceae Climax GC 5,89 0,58 _ 0,29 6,77 Tetrorchidium didymostemon (Baill.) Pax & Hoffm. Euphorbiaceae Pioneer GC 3,85 0,58 0,32 1,49 6,25 Holarrhena floribunda (G. DON.) Dur. & Schinz. Apocynaceae Pioneer GC-SZ 2,26 1,36 1,6 0,59 5,83 Erythrophleum suaveolens (Guill. & Pherr.) Brenan. Fabaceae Pioneer GC-SZ 2,26 _ 2,56 0,89 5,72 Myrianthus arboreus P. Beauv. Moraceae Pioneer GC 4,76 _ 0,64 0,29 5,7 Newbouldia laevis (P. Beauv.) Seemann. ex Bureau Bignoniaceae Pioneer GC 4,76 0,58 0,32 _ 5,66 Spondias monbin Linn. Anacardiaceae Pioneer GC-SZ 4,3 0,97 0,32 _ 5,6 Lecaniodiscus cupanioides Planch. & Benth. Sapindaceae Pioneer GC 4,98 0,19 0,32 _ 5,5 Spathodea campanulata P. Beauv. Bignoniaceae Climax GC 0,9 1,36 1,28 1,79 5,34 Trichilia heudelotii Planch. ex Oliv. Meliaceae Climax GC 3,17 0,97 0,96 _ 5,11 Harungana madagascariensis Lam. ex Poir. Guttiferae Pioneer GC 0,9 0,38 2,24 1,49 5,03 Vitex doniana Sweet Verbenaceae Pioneer GC-SZ 1,36 1,16 1,28 1,19 5 Celtis zenkeri Engl. Ulmaceae Climax GC 4,98 _ _ _ 4,98 Macaranga hurifolia Beille Euphorbiaceae Pioneer GC 4,08 0,38 0,32 _ 4,79 Dialium guineense Willd. Fabaceae Pioneer GC 2,94 _ 0,32 1,49 4,76 Rauvolfia vomitoria Afzel. Apocynaceae Pioneer GC-SZ 2,94 0,19 _ 1,19 4,33 Triplochiton scleroxylon K.Schum. Sterculiaceae Climax GC 3,4 0,38 _ 0,29 4,08 Uapaca guineensis Müll. Arg. Euphorbiaceae Climax GC 3,17 _ _ 0,89 4,07 Ricinodendron heudelotii (Baill.) Pierre ex Pax Euphorbiaceae Climax GC 1,36 1,94 0,32 0,29 3,92 Margaritaria discoidea (Baiil) Webster Phyllanthaceae Pioneer GC-SZ 0,22 0,38 0,64 2,38 3,64 140 J. Ecol. Nat. Environ.

Appendix S1. Contd.

Parinari glabra Oliv. Chrysobalanaceae Climax GC 3,4 _ _ _ 3,4 Aningeria altissima (A. Chev.) Aubrév. et Pellegr. Sapotaceae Climax GC 3,17 _ _ _ 3,17 Aphania senegalensis (Juss.ex Poir.) Radlk. Sapindaceae Climax GC 3,17 _ _ _ 3,17 Canthium schimperianum A. Rich. Rubiaceae Pioneer GC-SZ 3,17 _ _ _ 3,17 Piptadeniastrum africanum (Hook. f.) Brenan Fabaceae Climax GC 2,26 0,58 0,32 _ 3,17 Polyscias fulva (Hiern) Harms Araliaceae Climax GC 3,17 _ _ _ 3,17 Lonchocarpus sericeus (Poir.) Kunth Fabaceae Pioneer GC-SZ 1,13 1,36 _ 0,59 3,09 Hannoa klaineana Pierre et Engl. Simaroubaceae Climax GC 0,68 0,19 1,6 0,59 3,07 Macaranga heudelotii Baill. Euphorbiaceae Pioneer GC 1,58 1,16 0,32 _ 3,07 Nesogordonia papaverifolia (A.Chev.) R. Capuron Sterculiaceae Climax GC 2,72 _ 0,32 _ 3,04 Discoglypremna caloneura (Pax) Prain. Euphorbiaceae Pioneer GC 0,9 1,16 0,96 _ 3,03 Sterculia oblonga Mast. Sterculiaceae Climax GC 2,72 _ _ 0,29 3,01 Afzelia africana Sm. Fabaceae Pioneer GC-SZ 1,58 _ 0,64 0,59 2,82 Baphia nitida Lodd. Fabaceae Climax GC 2,72 _ _ _ 2,72 Bombax buonopozense P. Beauv. Bombacaceae Climax GC 0,22 0,97 0,32 1,19 2,71 Zanthoxylum macrophylla Engl. Rutaceae Climax GC 0,68 _ 1,92 _ 2,6 Holoptelea grandis (Hutch.) Mildbr. Ulmaceae Climax GC 2,26 0,19 _ _ 2,46 Dacryodes klaineana (Pierre) H.J.Lam Burseraceae Climax GC 2,04 _ _ 0,29 2,33 Manilkara multinervis (Baker) Dubard. Sapotaceae Pioneer GC-SZ 2,04 _ _ 0,29 2,33 Cathormion altissimum Hook.f.)Hutch. & Dandy Fabaceae Climax GC 2,26 _ _ _ 2,26 Pentaclethra macrophylla Benth. Fabaceae Climax GC 2,26 _ _ _ 2,26 Turraeanthus africanus Pellegr. Meliaceae Climax GC 2,26 _ _ _ 2,26 Pterygota macrocarpa K. Schum. Sterculiaceae Climax GC 1,81 _ 0,32 _ 2,13 Distemonanthus benthamianus Baill. Fabaceae Climax GC 0,9 0,19 0,96 _ 2,06 Maesopsis eminii Engl. Rhamnaceae Climax GC 2,04 _ _ _ 2,04 Mitragyna stipulosa O. Kuntze Rubiaceae Pioneer GC-SZ 2,04 _ _ _ 2,04 Ficus ovata Vahl. Moraceae Pioneer GC 0,45 0,38 _ 1,19 2,03 Blighia sapida Konig Sapindaceae Pioneer GC 0,22 _ 0,96 0,59 1,78 Canthium subcordatum DC. Rubiaceae Pioneer GC 1,58 0,19 _ _ 1,78 Berlinia grandiflora (Valh) Hutch. & Dalz. Fabaceae Pioneer GC-SZ 1,13 _ _ 0,59 1,73 Spondiathus preussii Engl. Euphorbiaceae Climax GC 1,36 _ _ 0,29 1,65 Eriocoelum kerstingii Gilg. & Engl. Sapindaceae Pioneer SZ 1,58 _ _ _ 1,58 Lovoa trichilioides Harms Meliaceae Climax GC 1,58 _ _ _ 1,58 Sorindea juglandifolia (A.Rich.)Planch.ex Oliv. Anacardiaceae Pioneer GC-SZ 1,58 _ _ _ 1,58 Vitex ferruginea Schum & Thonn. Verbenaceae Climax GC 1,58 _ _ _ 1,58 Amphimas pterocarpoïdes Harms Fabaceae Climax GC 0,22 0,38 0,64 0,29 1,55 Trichilia prieuriana A. Juss.subsp. prieuriana Meliaceae Climax GC 1,36 0,19 _ _ 1,55 Millettia zechiana Harms Fabaceae Pioneer GC 0,45 0,19 _ 0,89 1,54 Trichilia megalantha Harms Meliaceae Climax GC 1,13 0,38 _ _ 1,52 Tetrapleura tetraptera (Schum.& Thonn.) Taub. Fabaceae Climax GC 1,13 _ 0,32 _ 1,45 Khaya anthotheca (Welw.) C.DC. Meliaceae Climax GC 0,45 _ 0,32 0,59 1,37 Afzelia bella Harms var.gracilior Keay Fabaceae Climax GC 1,36 _ _ _ 1,36 Anogeissus leiocarpus (DC.) Guill. & Perr. Combretaceae Pioneer SZ 1,36 _ _ _ 1,36 Olax subscorpioidea Oliv. Olacaceae Pioneer GC 1,36 _ _ _ 1,36 Morus mesozygia Stapf Moraceae Climax GC 0,68 _ 0,32 0,29 1,29 Ficus populifolia Vahl Moraceae Pioneer SZ _ _ 1,28 _ 1,28 Albizia ferruginea (Guill.& Perr.)Benth. Fabaceae Climax GC _ 0,19 0,96 _ 1,15 Cassia sieberiana DC. Fabaceae Pioneer GC-SZ 0,22 _ 0,32 0,59 1,14 Dichapetalum oblongum (Hook.f.ex Benth.) Engel. Dichapetalaceae Climax GC 1,13 _ _ _ 1,13 Gaertneria paniculata Benth. Rubiaceae Pioneer GC 1,13 _ _ _ 1,13 Hexalobus crispiflorus A. Rich. Annonaceae Pioneer GC-SZ 1,13 _ _ _ 1,13 Mareya micrantha (Benth.) Muell. Euphorbiaceae Climax GC 1,13 _ _ _ 1,13 Adjossou et al. 141

Appendix S1. Contd.

Pachystela brevipes (Bak) Baill.ex Engl. Sapotaceae Pioneer GC 1,13 _ _ _ 1,13 Peddiea fischeri Engl. Thymelaeaceae Climax GC 1,13 _ _ _ 1,13 Pentadesma butyraccea Sabine Guttiferae Pioneer GC-SZ 1,13 _ _ _ 1,13 Syzygium guineense (Willd.) DC. Myrtaceae Pioneer SZ 1,13 _ _ _ 1,13 Klainedoxa gabonensis Pierre ex Engl. Irvingiaceae Climax GC 0,68 0,38 _ _ 1,07 Sterculia rhinopetala K. Schum. Sterculiaceae Climax GC 0,68 0,38 _ _ 1,07 Treculia africana Decne Moraceae Climax GC 0,45 _ _ 0,59 1,05 Parinari excelsa Sabine Chrysobalanaceae Climax GC 0,68 _ 0,32 _ 1 Sapium ellipticum (Hochst) Pax Euphorbiaceae Pioneer GC-SZ 0,68 _ 0,32 _ 1 Macaranga spinosa Müll. Arg. Euphorbiaceae Pioneer GC 0,68 _ _ 0,29 0,97 Dracaena arborea Link. Dracaenaceae Pioneer GC 0,22 0,38 _ 0,29 0,91 Gymnostemon zaizou Aubrév. & Pellegr. Simaroubaceae Climax GC 0,9 _ _ _ 0,9 Homalium letestui Pellegr Flacourtiaceae Climax GC 0,9 _ _ _ 0,9 Phoenix reclinata Jacq. Palmae Pioneer GC-SZ 0,9 _ _ _ 0,9 Canthium horizontale (Schum. & Thonn.) Hiern Rubiaceae Pioneer GC-SZ 0,68 0,19 _ _ 0,87 Markhamia tomentosa (Benth.) K.Schum. Bignoniaceae Pioneer GC 0,68 0,19 _ _ 0,87 Detarium senegalense J.F.Gmi Fabaceae Climax GC _ 0,19 0,32 0,29 0,81 Afrosersalisia afzelii (Engel.) A. Chev. Sapotaceae Climax GC 0,45 _ 0,32 _ 0,77 Bridelia atroviridis Müll. Arg. Euphorbiaceae Pioneer GC 0,45 _ _ 0,29 0,75 Antidesma laciniatum Mül. Arg. var. membranaceum Müll. Arg. Euphorbiaceae Climax GC 0,68 _ _ _ 0,68 Ficus polita Vahl. Moraceae Pioneer GC-SZ 0,68 _ _ _ 0,68 Ekebergia senegalensis A. Juss. Meliaceae Pioneer SZ _ _ 0,64 _ 0,64 Blighia welwitschii (Hiern) Radlk Sapindaceae Climax GC 0,22 0,38 _ _ 0,61 Albizia coriaria Welw.ex Oliv. Fabaceae Pioneer GC-SZ _ _ _ 0,59 0,59 Trema orientalis (Linn.) Bl. Ulmaceae Pioneer GC-SZ _ _ _ 0,59 0,59 Nauclea diderrichii (de Wild. & Th. Dur. Merril.) Rubiaceae Climax GC _ 0,58 _ _ 0,58 Strombosia glauscescens J.Leonard var. lucida Olacaceae Climax GC _ 0,58 _ _ 0,58 Vernonia colorata (Willd.) Drake Compositae Pioneer GC-SZ _ 0,58 _ _ 0,58 Rothmannia longiflora Salisb. Rubiaceae Climax GC 0,22 _ 0,32 _ 0,54 Anthocleista nobilis G. Don Loganiaceae Pioneer GC 0,22 _ _ 0,29 0,52 Albizia glaberrima (Schum. & Thonn.) Benth. Fabaceae Climax GC _ 0,19 0,32 _ 0,51 Blighia unijugata Bak. Sapindaceae Pioneer GC _ 0,19 0,32 _ 0,51 Ficus lyrata Warb. Moraceae Pioneer GC _ 0,19 0,32 _ 0,51 Drypetes gilgiana(Pax)(Pax)&K.Hoffm. Euphorbiaceae Climax GC 0,45 _ _ _ 0,45 Entandrophragma cylindricum (Welw.) C.DC. Meliaceae Climax GC 0,45 _ _ _ 0,45 Ficus tesselata Warb. Moraceae Climax GC 0,45 _ _ _ 0,45 Ficus umbellata Vahl. Moraceae Pioneer GC-SZ 0,45 _ _ _ 0,45 Lannea nigritana (Sc. Elliot) Keay var.nigritana Anacardiaceae Pioneer GC-SZ 0,45 _ _ _ 0,45 Malacantha alnifolia (Bak.) Pierre Sapotaceae Pioneer GC-SZ 0,45 _ _ _ 0,45 Millettia thonningii (Schum. & Thonn. ) Bak. Fabaceae Pioneer GC 0,45 _ _ _ 0,45 Musanga cecropioides R.Br. ex Tedlie Cecropiaceae Pioneer GC 0,45 _ _ _ 0,45 Napoleonaea vogelii Hook. ex Planch. Lecythidaceae Pioneer GC 0,45 _ _ _ 0,45 Pancovia pedicellaris Radlk.& Gilg Sapindaceae Climax GC 0,45 _ _ _ 0,45 Rinorea yaundensis Engl. Violaceae Climax GC 0,45 _ _ _ 0,45 Rothmannia urcelliformis (Hiern) Bullock.ex Robyns Rubiaceae Climax GC 0,45 _ _ _ 0,45 Stereospermum acuminatissimum K. Schum. Bignoniaceae Climax GC 0,45 _ _ _ 0,45 Tarenna pavettoides (Harv.) Sim Rubiaceae Climax GC 0,45 _ _ _ 0,45 Uapaca heudelotii Baill. Euphorbiaceae Pioneer GC-SZ 0,45 _ _ _ 0,45 Mansonia altissima A. Chevalier Sterculiaceae Climax GC 0,22 0,19 _ _ 0,42 Dracaena mannii Bak. Dracaenaceae Pioneer GC _ 0,38 _ _ 0,38 Ficus lutea Vahl. Moraceae Pioneer GC _ _ 0,32 _ 0,32 Garcinia afzelii Engl. Guttiferae Pioneer GC-SZ _ _ 0,32 _ 0,32 142 J. Ecol. Nat. Environ.

Appendix S1. Contd.

Zanthoxylum leprieurii Guill. & Perr. Rutaceae Pioneer GC _ _ _ 0,29 0,29 Aidia genipiflora (D.C.) Dandy Rubiaceae Climax GC 0,22 _ _ _ 0,22 Bertiera racemosa (D. Don.) K. Schum. Rubiaceae Climax GC 0,22 _ _ _ 0,22 Cordia platythyrsa Baker Boraginaceae Climax GC 0,22 _ _ _ 0,22 Cordia senegalensis Juss. Boraginaceae Climax GC 0,22 _ _ _ 0,22 Dichrostachys cinerea (L.) Wight & Arn. Fabaceae Pioneer SZ 0,22 _ _ _ 0,22 Diospyros mespiliformis Hochst. Ex A. DC. Ebenaceae Pioneer GC-SZ 0,22 _ _ _ 0,22 Dovyalis zenkeri Gilg. I.c Flacourtiaceae Pioneer GC 0,22 _ _ _ 0,22 Drypetes aframensis Hutch. Euphorbiaceae Climax GC 0,22 _ _ _ 0,22 Drypetes aylmeri Hutct .& Dalz. Euphorbiaceae Climax GC 0,22 _ _ _ 0,22 Drypetes leonensis Pax Euphorbiaceae Climax GC 0,22 _ _ _ 0,22 Erythrina vogelii Hook. f. Fabaceae Climax GC 0,22 _ _ _ 0,22 Ficus thonningii Blume Moraceae Pioneer GC-SZ 0,22 _ _ _ 0,22 Flacourtia flavescens Willd. Flacourtiaceae Pioneer SZ 0,22 _ _ _ 0,22 Garcinia ovalifolia Oliv. Guttiferae Pioneer GC-SZ 0,22 _ _ _ 0,22 Grewia pubescens P.Beauv. Tiliaceae Pioneer SZ 0,22 _ _ _ 0,22 Irvingia robur Mildbr. Irvingiaceae Climax GC 0,22 _ _ _ 0,22 Mimusops kummel Hochst. Sapotaceae Pioneer SZ 0,22 _ _ _ 0,22 Morelia senegalensis A.Rich.ex DC Rubiaceae Pioneer GC-SZ 0,22 _ _ _ 0,22 Parkia bicolor A.Chev. Fabaceae Climax GC 0,22 _ _ _ 0,22 Parkia filicoidea Welw. ex Oliv. Fabaceae Climax GC 0,22 _ _ _ 0,22 Psychotria psychotrioides (DC.) Roberty Rubiaceae Pioneer GC-SZ 0,22 _ _ _ 0,22 Pterygota bequaertii De Wild. Sterculiaceae Climax GC 0,22 _ _ _ 0,22 Rhodognaphalon brevicuspe (Sprague) Roberty Sterculiaceae Climax GC 0,22 _ _ _ 0,22 Symphonia globulifera Linn. f. Guttiferae Climax GC 0,22 _ _ _ 0,22 Voacanga africana Stapf. Apocynaceae Pioneer GC 0,22 _ _ _ 0,22 Zanha golungensis (Hiern) Sapindaceae Pioneer GC-SZ 0,22 _ _ _ 0,22 Erythrina mildbraedii Harms Fabaceae Climax GC _ 0,19 _ _ 0,19 Ficus vogeliana (Miq.) Miq. Moraceae Pioneer GC _ 0,19 _ _ 0,19 Monodora tenuifolia Benth. Annonaceae Pioneer GC _ 0,19 _ _ 0,19 Vernonia conferta Benth. Compositae Pioneer GC _ 0,19 _ _ 0,19

FR: Forest Relics; COCS: Coco System; COFS: Coffee System; FS: Follow Systems. GC-SZ: species encountered in several phytochoria in continental tropical Africa, GC: Guinean-Congolese regional centre of endemism species, mainly consists of humid forest domains, SZ: Sudanese-zambezian regional centre of endemism , mainly consists of savannah or dry vegetation specie.

Pioneer : First stage of natural succession ; Climax : last stage of natural succession